Journal of Forest Research

, Volume 18, Issue 1, pp 121–127 | Cite as

Does sika deer overabundance exert cascading effects on the raccoon dog population?

  • Yoshikazu SekiEmail author
  • Masaaki Koganezawa
Original Article


Habitat structure in Oku-Nikko, Japan, has been substantially modified by the overabundance of sika deer (Cervus nippon). A deer-proof fence (15.0 km and 900 ha) was constructed in 2001 to conserve vegetation. Although the understory inside the fence is dominated by Sasa nipponica (hereafter, Sasa), an important forage plant for deer, that outside the fence is dominated by Aster ageratoides leiophyllus (hereafter, Aster), an unpalatable plant to deer, and, partly, by bare floor. In this study, we examined the effects of deer on ground-dwelling insects and earthworms, the primary food resources of raccoon dogs (Nyctereutes procyonoides), and, thus, the bottom-up cascading effects of the herbivore on the omnivorous carnivore. Between July and September 2008, we examined the abundance of insects and earthworms by pitfall trapping and hand-sorting methods, respectively, both inside and outside the fence. The abundance of earthworms and insects (Scarabaeidae and Rhaphidophoridae) was higher on forest floors with Aster and/or bare floors outside the fence than on those with Sasa inside the fence. These results indicate that the increasing deer population in this area probably increased the number of these invertebrates outside the fence by modifying understory vegetation and/or depositing dung. Furthermore, the sighting rates of raccoon dogs obtained by spotlight counts were greater outside than inside the fence, suggesting that deer probably exert bottom-up cascading effects on raccoon dogs, at least during May to November, when the invertebrates are predominantly fed on by the omnivorous carnivore.


Bottom-up trophic cascade Higher trophic level Indirect effect Invertebrate Omnivorous carnivore 



We are grateful to Dr Tatsuhiro Okubo, Utsunomiya University, Dr Koichi Kaji and Dr Yayoi Kaneko, Tokyo University of Agriculture and Technology, Dr Takeshi Yasue, Ibaraki University, Ms Reiko Horie, Utsunomiya University, and two anonymous referees for their valuable advice and constructive criticism of the manuscript. In this study we used the data on spotlight counts conducted from 2002 to 2010 by the students of the Wildlife Management Laboratory of Utsunomiya University. We are grateful to them for the use of the data.


  1. Asada M, Ochiai K (1996) Food habits of sika deer on the Boso Peninsula, central Japan. Ecol Res 11:89–95CrossRefGoogle Scholar
  2. Bardgett RD, Wardle DA (2003) Herbivore-mediated linkages between aboveground and belowground communities. Ecology 84:2258–2268CrossRefGoogle Scholar
  3. Campos-Arceiz A, Takatsuki S (2005) Food habits of sika deer in the Shiranuka Hills, eastern Hokkaido: a northern example from the north-south variations in food habits in sika deer. Ecol Res 20:129–133CrossRefGoogle Scholar
  4. Cote SD, Rooney TP, Tremblay JP, Dussault C, Waller DM (2004) Ecological impacts of deer overabundance. Annu Rev Ecol Evol Syst 35:113–147CrossRefGoogle Scholar
  5. Dyer LA, Letourneau DK (1999) Relative strengths of top-down and bottom-up forces in a tropical forest community. Oecologia 119:265–274CrossRefGoogle Scholar
  6. Flowerdew JR, Ellwood SA (2001) Impacts of woodland deer on small mammal ecology. Forestry 74:277–287CrossRefGoogle Scholar
  7. Hasegawa J (2008) Changes in nature of Tochigi Prefecture. Self publication, Tochigi (in Japanese)Google Scholar
  8. Hunter MD, Price PW (1992) Playing chutes and ladders: heterogeneity and the relative roles of bottom-up and top-down forces in natural communities. Ecology 73:724–732Google Scholar
  9. Jayasekara P, Takatsuki S (2000) Seasonal food habits of sika deer population in the warm temperature forests of the westernmost part of Honshu, Japan. Ecol Res 15:153–157CrossRefGoogle Scholar
  10. Kagata H, Ohgushi T (2006) Bottom-up trophic cascades and material transfer in terrestrial food webs. Ecol Res 21:26–34CrossRefGoogle Scholar
  11. Kaneko Y, Maruyama N, Macdonald DW (2006) Food habits and habitat selection of suburban badgers (Meles meles) in Japan. J Zool 20:78–89Google Scholar
  12. Kanzaki N, Maruyama N, Koganezawa M, Taniguchi M (1998) Tree barking by sika deer in Nikko, Tochigi prefecture. Wildl Conserv Jpn 3:107–117 (in Japanese with English summary)Google Scholar
  13. Koganezawa M (1983) Altitudinal distribution of middle-sized mammals in Nikko area, Tochigi Prefecture (I). Mem Tochigi Pref Mus 1:39–66 (in Japanese with English summary)Google Scholar
  14. Koganezawa M, Kurokawa M (1983) Altitudinal distribution of middle-sized mammals in Nikko area, Tochigi Prefecture (II)—focused on the winter distribution and the food habits of the red fox (Vulpes vulpes). Mem Tochigi Pref Mus 1:67–82 (in Japanese with English summary)Google Scholar
  15. Koganezawa M, Li Y (2002) Sika deer response to spotlight counts: implications for distance sampling of population density. Mammal Study 27:95–99CrossRefGoogle Scholar
  16. Koganezawa M, Satake C (1996) Effects of grazing by sika-deer on the vegetation of Oku-Nikko and their management. Trans Nat Found Proj 5:57–66 (in Japanese with English summary)Google Scholar
  17. Koike S, Kato M, Morimoto H, Furubayashi K (2006) Study of dung beetles in feces of sika deer, Cervus nippon, in Tanzawa Mountains, central Japan. Wildl Conserv Jpn 10:45–60 (in Japanese with English summary)Google Scholar
  18. Koike S, Morimoto H, Goto Y, Kozakai C, Yamazaki K (2008) Frugivory of carnivores and seed dispersal of fleshy fruits in cool-temperate deciduous forests. J For Res 13:215–222CrossRefGoogle Scholar
  19. Kondo T (1980) Food habits of the red fox (Vulpes vulpes japonica) and the Japanese marten (Martes melampus melampus). Bull Osaka Kyoiku Univ 29:19–23 (in Japanese with English summary)Google Scholar
  20. McCauley DJ, Keesing F, Young TP, Allan BF, Pringle RM (2006) Indirect effects of large herbivores on snakes in an African Savanna. Ecology 87:2657–2663PubMedCrossRefGoogle Scholar
  21. Melis C, Buset A, Aarrestad PA, Hanssen O, Meisingset EL, Andersen R, Moksnes A, Roskaft E (2006) Impact of red deer Cervus elaphus grazing on bilberry Vaccinium myrtillus and composition of ground beetle (Coleoptera, Carabidae) assemblage. Biodivers Conserv 15:2049–2059CrossRefGoogle Scholar
  22. Melis C, Sundby M, Andersen R, Moksnes A, Pedersen B, Roskaft E (2007) The role of moose Alces alces L. in boreal forest: the effect on ground beetles (Coleoptera, Carabidae) abundance and diversity. Biodivers Conserv 16:1321–1335CrossRefGoogle Scholar
  23. Morin PJ (1999) Community ecology. Blackwell, OxfordGoogle Scholar
  24. Nagata Y, Yoshida M (1997) The basis of multiple comparison procedure. Scientist, Tokyo (in Japanese)Google Scholar
  25. Pace ML, Cole JJ, Carpenter SR, Kitchell JF (1999) Trophic cascades revealed in diverse ecosystems. Trends Ecol Evol 14:483–488PubMedCrossRefGoogle Scholar
  26. Persson L (1999) Trophic cascades: abiding heterogeneity and the trophic level concept at the end of the road. Oikos 85:385–397CrossRefGoogle Scholar
  27. R Development Core Team (2009) R: a language and environment for statistical computing. R Foundation for Statistical Computing, Vienna, Austria.
  28. Rambo JL, Faeth SH (1999) Effects of vertebrate grazing on plant and insect community structure. Conserv Biol 13:1047–1054CrossRefGoogle Scholar
  29. Rooney TP, Waller DM (2003) Direct and indirect effects of white-tailed deer in forest ecosystems. For Ecol Manag 181:165–176CrossRefGoogle Scholar
  30. Sasaki H, Kawabata M (1994) Food habits of the raccoon dog Nyctereutes procyonoides viverrinus in a mountainous area of Japan. J Mammal Soc Jpn 19:1–8Google Scholar
  31. Schmitz OJ, Hamback PA, Beckrman AP (2000) Trophic cascades in terrestrial systems: a review of the effects of carnivore removals on plants. Am Nat 155:141–153PubMedCrossRefGoogle Scholar
  32. Seki Y (2011) Trophic interaction between the sika deer and the raccoon dog in Oku-Nikko, Japan. PhD thesis, Tokyo University of Agriculture and Technology (in Japanese)Google Scholar
  33. Seki Y, Koganezawa M (2010) Factors influencing the increase in earthworms outside deer-proof fences in Oku-Nikko, central Japan: the influence of the modification of understory vegetation by sika deer. J Jpn For Soc 92:241–246 (in Japanese with English summary)CrossRefGoogle Scholar
  34. Seki Y, Koganezawa M (2011) Factors influencing winter home ranges and activity patterns of raccoon dogs Nyctereutes procyonoides in a high-altitude area of Japan. Acta Theriol 56:171–177CrossRefGoogle Scholar
  35. Shibata E, Hino T (2009) Ecology of sika deer and forest ecosystem of Mt. Ohdaigahara. Tokai University Press, Kanagawa (in Japanese)Google Scholar
  36. Stewart AJA (2001) The impact of deer on lowland woodland invertebrates: a review of the evidence and priorities for future research. Forestry 74:259–270CrossRefGoogle Scholar
  37. Strong DR (1992) Are trophic cascades all wet? Differentiation and donor-control in speciose ecosystems. Ecology 73:747–754CrossRefGoogle Scholar
  38. Suominen O, Danell K, Bergstrom R (1999a) Moose, trees, and ground-living invertebrates: indirect interactions in Swedish pine forests. Oikos 84:215–226CrossRefGoogle Scholar
  39. Suominen O, Danell K, Bryant JP (1999b) Indirect effects of mammalian browsers on vegetation and ground-dwelling insects in an Alaskan floodplain. Ecoscience 6:505–510Google Scholar
  40. Suominen O, Niemela J, Martikainen P, Niemela P, Kojola I (2003) Impact of reindeer grazing on ground-dwelling Carabidae and Curculionidae assemblages in Lapland. Ecography 26:503–513CrossRefGoogle Scholar
  41. Takahashi H, Kaji K (2001) Fallen leaves and unpalatable plants as alternative foods for sika deer under food limitation. Ecol Res 16:257–262CrossRefGoogle Scholar
  42. Takatsuki S (1983) The importance of Sasa nipponica as a forage for sika deer (Cervus nippon) in Omote-Nikko. Jpn J Ecol 33:17–25Google Scholar
  43. Takatsuki S (1986) Food habits of sika deer on Mt. Goyo, northern Honshu. Ecol Res 1:119–128CrossRefGoogle Scholar
  44. Takatsuki S (2009) Effects of sika deer on vegetation in Japan: a review. Biol Conserv 142:1922–1929CrossRefGoogle Scholar
  45. Ueda A, Hino T, Ito H (2009) Relationships between browsing on dwarf bamboo (Sasa nipponica) by sika deer and the structure of ground beetle (Coleoptera: Carabidae) assemblage. J Jpn For Soc 91:111–119 (in Japanese with English summary)CrossRefGoogle Scholar
  46. Wardle DA, Barker GM, Yeates GW, Bonner KI, Ghani A (2001) Introduced browsing mammals in New Zealand natural forests: aboveground and belowground consequences. Ecol Monogr 71:587–614CrossRefGoogle Scholar
  47. Yamamoto Y (1991) Food habits of Meles meles anakuma in Mt. Nyugasa, Nagano Pref., Japan. Nat Environ Sci Res 4:73–83 (in Japanese with English summary)Google Scholar
  48. Yamamoto Y (1994) Comparative analyses on food habits of Japanese marten, red fox, badger and raccoon dog in Mt. Nyugasa, Nagano Prefecture, Japan. Nat Environ Sci Res 7:45–52 (in Japanese with English summary)Google Scholar
  49. Yokoyama S, Shibata E (1998) Characteristics of Sasa nipponica grassland as a summer forage resource for sika deer on Mt. Ohdaigahara, central Japan. Ecol Res 13:193–198CrossRefGoogle Scholar
  50. Yokoyama S, Koizumi T, Shibata E (1996) Food habits of sika deer as assessed by fecal analysis in Mt. Ohdaigahara, central Japan. J For Res 1:161–164CrossRefGoogle Scholar
  51. Yokoyama M, Kaji K, Suzuki M (2000) Food habits of sika deer and nutritional value of sika deer diets in eastern Hokkaido, Japan. Ecol Res 15:345–355CrossRefGoogle Scholar

Copyright information

© The Japanese Forest Society and Springer 2012

Authors and Affiliations

  1. 1.United Graduate School of Agricultural ScienceTokyo University of Agriculture and TechnologyTokyoJapan
  2. 2.Utsunomiya University Forests, Faculty of AgricultureUtsunomiya UniversityShioyaJapan

Personalised recommendations